Physiological and Morphological Basis for Differences in Growth, Water Use and Drought Resistance Among Cercis L. Taxa Dissertat
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PHYSIOLOGICAL AND MORPHOLOGICAL BASIS FOR DIFFERENCES IN GROWTH, WATER USE AND DROUGHT RESISTANCE AMONG CERCIS L. TAXA DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Petra Sternberg, M.S. Graduate Program in Horticulture and Crop Science The Ohio State University 2012 Dissertation Committee: Daniel K. Struve, Advisor David J. Barker James D. Metzger Daniel A. Herms Copyrighted by Petra Sternberg 2012 Abstract Water is one of the principle factors determining the structure and species composition of the vegetation occupying a given site. Plantings in constructed sites are particularly problematic as urban environments magnify stresses that are common to most landscapes. Often, trees are selected predominantly for their aesthetic contribution to the landscape, with little consideration of their adaptation to the site. Selecting trees with improved drought resistance and reduced water use may be the best strategy to improve survival, growth and health of trees in urban and suburban landscapes. However, the variety of plant characteristics that contribute to drought resistance and the variation in site characteristics makes plant selection for dry sites difficult. Cercis are small to medium sized and often multi-stemmed trees. Cercis canadensis L. is a valuable commercially- produced landscape tree in the United States. The aim of this study was to investigate the inter-taxa variation in drought resistance and water use within the genus Cercis, thereby providing information as to their usefulness for plantings in urban landscapes and to facilitate breeding for improved water use and drought resistance. The genus Cercis currently includes ten recognized species native to the warm, north- temperate zones of North America and Eurasia. The exact number of species and their delimitation remains controversial. ii The first objective was to describe the Cercis taxa to be used in subsequent experiments (Chapters 3 and 4), to compare them to Cercis described in the literature and describe morphological differences that can influence drought resistance. Growth habit and leaf characteristics were measured and principal component analysis employed to describe variation in the observed characteristics. The taxa described were representative of their species and/or variety. Cercis taxa differed widely in their growth habit and leaf characteristics and showed adaption to their native environment. Cercis employed common as well as taxa-specific strategies to deal with similar environmental conditions. The persistence of the leaf characteristics under non-limiting soil moisture conditions indicated that the characteristics typical of the taxa grown in their native environment were under genetic control. The observation that some characteristics on the Eastern North American taxa espressed great variability indicated introgression of these charateristics and that the taxa are still in the process of speciation. The root system needs to enable plants to meet their transpirational demand and is a key component of plant adaption to xeric environments. Thus, the second objective was to evaluate water use characteristics and morphology of seven Cercis L. taxa of different origins and to determine the growth strategies of the taxa in terms above versus below ground biomass under non-limiting soil moisture conditions. Seven Cercis L. taxa were grown from seed in the greenhouse, and 60 days after germination, water use plant-1 day-1 was determined gravimetrically, three times over 72 h. Height, caliper, number of nodes on the main shoot axis and leaf area were measured. Roots were scanned and analyzed with WinRhizo software to determine root length, root surface area and root diameter. iii Dry weight of leaves, shoots and roots was obtained. A principal component analysis showed great differences in shoot and root morphology as well as water use characteristics among taxa. It showed a trend towards decreasing water use per unit plant mass with increasing plant size. No consistent pattern of characteristics employed by mesic- or xeric-adapted taxa could be found. Each taxon uses a unique set of characteristics to adapt to its original environment. Plants have developed morphological, anatomical and physiological adaptations to resist drought. The third objective was to determine morphological and physiological differences among Cercis taxa in response to water deficit. Seedlings of seven Cercis taxa were submitted to a drought and recovery cycle under greenhouse conditions. Net photosynthesis, stomatal conductance, intrinsic water use efficiency and chlorophyll fluorescence, as well as plant growth and water use were measured. Paraheliotropism was observed in all taxa at varying degrees. All taxa reduced net photosynthesis and stomatal conductance shortly after exposure to limited water availability. Cercis occidentalis and C. siliquastrum presented with high initial values for net photosynthesis and stomatal conductance and showed a relatively lower decline in both values than other taxa. Chlorophyll fluorescence declined in some taxa during the drought period, while other taxa where unaffected. Affected taxa showed inclining values for net photosynthesis, stomatal conductance and chlorophyll fluorescence during the recovery period. Taxa in this study are drought-resistant and employ several common drought resistant mechanisms. One is the ability of plants to withstand drought stress without altering the function of the PSII system and/or the capacity of plants to recover from damage another is paraheliotropism. Overall, the Cercis taxa displayed great inter-taxa variation. Some iv common drought resistance mechanisms were found, however, other mechanisms, especially differences in growth habit and morphology appeared to be more specific adaptations to the native environment of each taxon; for instance a shrub-like growth habit, consistent with capturing precipitation via stem flow, and/or glaucousness of the leaves. Cercis canadensis var. texensis would be an excellent candidate for inducing drought resistance into a breeding program based on the combination of relatively high drought resistance displayed in these experiments, its upright growth habit and rapid growth rate. v Dedication I dedicate this dissertation to my loving husband Kurt Bresko and my adorable son Christopher (“Muckel”). Thank you for your smiles! vi Acknowledgments I wish to thank everybody whose assistance, advice and support have made the completion of my graduate studies and this dissertation possible. First, I would like to express my sincere gratitude to my doctoral advisor Dr. Daniel Struve for his guidance and advice. I would also like to thank my graduate committee, Dr. David Barker, Dr. James Metzger and Dr. Daniel Herms, for their direction, dedication and invaluable advice along this project. I am indebted to my parents for their love and emotional support. I am especially thankful to my son, Christopher Bresko, for being the ultimate reason for finishing this dissertation, and to my husband Kurt Bresko, for his love and encouragement. Without this invaluable team the completion of my dissertation would have not been possible. vii Vita 2003 ............................................................ Diplom-Ingenieur Gartenbau (FH), University of Applied Science Osnabrück, Germany 2007 ............................................................ M.S. Horticulture and Crop Sciences, The Ohio State University 2007 to present ............................................ Graduate Research and Teaching Associate, Department of Horticulture and Crop Science, The Ohio State University Publications Sternberg, P. and D.K. Struve, 2008 Cyclanilide foliar applications induce grater lateral branching than pruning in container grown whips. Journal of Environmental Horticulture 26(1): 45-50 Sternberg, P. and D.K. Struve, 2007 Cyclanilide spray increases branching in containerized whip production. Journal of Environmental Horticulture 25(4): 221-228 Struve, D.K., Sternberg, P. , Drunasky, N., Bresko, K.L. and R. Gonzales, 2006 Growth and water use characteristics of six Eastern North American Oak (Quercus) species and the implications for urban forestry. Arboriculture and Urban Forestry 32: 202-213 Fields of Study Major Field: Horticulture and Crop Science viii Table of Contents Abstract .......................................................................................................................... ii Dedication ..................................................................................................................... vi Acknowledgments ........................................................................................................ vii Vita ............................................................................................................................. viii List of Tables................................................................................................................ xii List of Figures ..............................................................................................................xiv CHAPTER 1 INTRODUCTION ...........................................................................................................1 1.1 Statement of the problem and significance ..............................................................2 1.2 Description and taxonomy of the genus Cercis L. ...................................................4